Overview

Brief Summary

Species Abstract

The Painted Hunting Dog is one of the most critically endangered carnivores in Africa, with approximately 5000 individuals extant. The early Holocene species population is estimated at about one half million, and the prehistoric range covered virtually all of the non-desert, non-jungle area of sub-Saharan Africa. In the last several decades, the habitat of L. Pictus has become severely fragmented due to expansion of the human population in Africa, and numbers have dwindled. The species has been historically termed "African Wild Dog", which name is declining in usage, due to pejorative aspects of such a name.

There are five acknowledged subspecies: L.p. pictus (Southern Africa), L.p. lupinus (East Africa), L.p. manguensis (West and Central Africa), L.p. saharicus (Sahara) and L.p. somalicus (Horn of Africa). Based upon the detailed geographic analysis in the following, the last three subspecies can be clearly considered critically endangered due to the extremely low populations in their sub-regions and the ongoing severe threats in those areas.

Until as recently as the 1970s L. pictus had a broad distribution within sub-Saharan Africa. Presently Botswana has one of the most robust populations with an estimated 150 individuals in the northeast part of the Okavango Delta, encompassing the eastern part of the Moremi Game Reserve. The area is approximately 3,000 square kilometres. This region is protected not just by paper legislation, but due to the care and concern of the Botswanan people. In Namibia, approximately 200 wild dogs are estimated to have existed by 2008, with the population dying off by about ten percent per annum. Virtually all of the sightings are in Tsumkwe East and West Caprivi. Livestock grazing and poaching account for the majority of the decline of the species in Namibia.

There is little sexual dimorphism in this species with adults attaining body mass to 36 kg and lengths to 110 cm plus tail extension to an additional 40 cm.(Novak) Pelage exhibits a short fur of mottled yellow, red, black and white patches. This animal is lean and muscular, built for running endurance. Each foot manifests four toes. Characteristically the bushy tail has a white tip. Ears are ovoid and sizable.

The pack chooses a den location which affords a good burrow, but which also ideally has some shade to protect pups from the harsh sun when they first begin to venture from the den. This shade also helps to mitigate against high burrow temperatures in the hot season. A typical burrow landform might consist of loamy soil which was colonised by termites. After construction of the termite mound to pulverise and soften the soil, an aardvark may have used this locale for rooting. A culmination of decades of earthwork by these two pioneer species will then be the formation of a more intricate den by the Painted Hunting Dog pack, building upon the previous subsurface work of others.

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Description

The African wild dog is one of the world's most social and distinctive canids. The short, wiry coat is coloured in blotches of yellow, grey, black and white (2) that give rise to the scientific name of Lycaon pictus or 'painted wolf-like animal' in Greek (3). The hair is short on the limbs and body and longer on the neck. Each dog has a unique colouration pattern, and this is used by researchers to identify individuals (2). The body is thin and muscular, the tail bushy with a white tip and the legs are long (2) (4). Males are slightly larger than females (2). Unlike the other canid species there are only four, rather than five, toes on the front feet (2) (4). These dogs have large rounded ears, which probably help with heat loss as well as keeping track of pack members by picking up long distance vocal signals (2) (4).

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Biology

African wild dogs are highly sociable and exhibit a very unusual social system; within their packs, dogs of the same sex are closely related to each other but not to individuals of the opposite sex, and only the dominant male and female will breed (4) (5) (6). Packs vary in size from 2 to 27 individuals and new packs are formed when subgroups of the same sex (usually siblings) disperse and join up with a subgroup of the opposite sex (5), leading to the unusual configuration of the pack. Only the dominant female will give birth to pups, and births may take place throughout the year, although they are more common between March and June (2). Litter size is the largest of any canid, averaging ten pups (2); these are born within dens where they remain for around three months (6). Initially the mother will stay with her pups and when members of the pack return from hunting they regurgitate food for her. As the pups get older however, all pack members help with feeding and 'baby sitting' of the young dogs (6). Juveniles are fully independent at 16 to 24 months (6) but remain with their pack, females are more likely to disperse, usually leaving in a sub-group with their sisters once they reach two years old (6). Outside of the breeding season, wild dogs are nomadic and wander over large distances in search of prey; home ranges can be as large as 5,000 square kilometres, but are often restricted to areas of less than 200 square kilometres (2). These dogs are carnivorous and hunt their prey by cooperating closely in a group (2). This strategy enables them to hunt prey comprising antelope and ungulates much larger than themselves, to include kudu bulls and wildebeest weighing up to 250 kilograms, as well as ensuring their hunting success is much higher than that of other large carnivorous species (2). Packs set out to hunt in the cool of dawn and dusk, avoiding other predators such as lions. The victim is pulled to the ground and the group descends to feed; pups in the pack are allowed to eat first (6).

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The tri-coloured African wild dog (Lycaon pictus) is probably best known for its organised and well-structured social system (Girman et al., 1997). Pack sizes are based on a trade-off of energetic costs and benefits (Hayward et al., 2006), with larger packs (approx. 20-40 adults) experiencing fitness benefits in terms of hunting success and pup survival (Woodroffe, 2011). Their social structure allows for a dominant breeding pair, with a single breeding female, but rearing of the pups becomes the responsibility of the sub-dominant “helpers” in the pack (Courchamp and Macdonald, 2001).

These mammals are medium sized carnivores (20-28 kg) and mainly prey on hoofed animals (ungulates), such as eland and zebra (although it has been reported that they can bring down juvenile buffalo), hunting at dusk and dawn and consuming more meat per body weight than any other carnivore (Hayward et al., 2006). They typically inhabit open woodland, where competition pressure from other predators such as lions, is slightly reduced (Vucetich and Creel, 2001).

Due to their large range-size requirements, inevitable conflict with humans has been, and continues to be, the primary reason for the decline in numbers of this endangered mammal (IUCN Red List of Threatened Species). Excessive hunting several decades ago drastically reduced the population size of the African wild dogs, now occupying less than 7% of their original geographic range (Woodroffe, 2011).

Conservation bodies such as The African Wildlife Foundation and the African Wild Dog Conservancy focus on monitoring pack sizes and activity, educating local communities and reducing poaching activity in order to stabilise current populations and boost those that are showing a decline. Additionally, the creation of corridors to link scattered populations and increase gene-flow between populations could prove invaluable for the on-going survival of these species (Davies-Mostert et al., 2012).

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MammalMAP: African Wild dogs

African Wild dogs or Lycaon pictus (stemming from the Greek word for ‘wolf’ and the latin word for ‘painted’) are the only canid species to lack dewclaws on their forelimbs. Africa’s largest canid – an adult weighs within the range of 18 – 36 kgs.

African Wild dogs form packs that on average consists of 7 – 15 members. Before the recent decline in the population, packs consisting of 100 members have been recorded. Animals within the pack have a unique social concerns and structure – from cooperation during hunts to regurgitating food to feed the young, sick, elderly or wounded.

Typically, wild dogs feed on ungulates that are often twice their body weight. The hunt is lead by the alpha male usually in the morning or early evening. Wild dogs locate prey by sight and pursue it in high speed chases that can reach 55 km/hr until the prey is exhausted.

The IUCN has listed African Wild dogs as an endangered species since 1996. The decline in population size has been attributed to shrinking habitats, conflict with livestock and game farmers, road accidents and infectious diseases.

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Distribution

Range Description

Historical data indicate that African Wild Dogs were formerly distributed throughout sub-Saharan Africa, from desert (Lhotse 1946) to mountain summits (Thesiger 1970), and probably were absent only from lowland rainforest and the driest desert (Schaller 1972). They have disappeared from much of their former range. The species is virtually eradicated from North and West Africa, and greatly reduced in Central Africa and North-east Africa. The largest populations remain in southern Africa (especially northern Botswana, western Zimbabwe, eastern Namibia, and western Zambia) and the southern part of East Africa (especially Tanzania and northern Mozambique).

The current geographic distribution of African Wild Dogs was estimated using data compiled by the IUCN SSC range-wide conservation planning process for Cheetahs and African Wild Dogs, including regional strategies (IUCN SSC 2008, in prep.) and subsequent associated national action plans (www.cheetahandwilddog.org). Current African Wild Dog range was considered to comprise only the resident range identified by participants in the IUCN SSC process: this represents land where participants were confident that African Wild Dogs had been confirmed to be resident within the previous 10 years. Land where residence was not confirmed (e.g., possible range, unknown range) was excluded.

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At one time the distribution of Lycaon pictus was throughout the non-forested and non-desert areas of Africa. Their current distribution is more fragmented. African hunting dogs are now found in Namibia, Botswana, Mozambique, parts of Zimbabwe, Swaziland, and the Transvaal.

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Range

The African wild dog was formerly present throughout sub-Saharan Africa, although it was never locally abundant (2) (4). This species is today restricted to fragmented populations mainly in southern and eastern Africa (2) (4). Potentially viable populations currently exist in Botswana, Kenya, Mozambique, South Africa, Tanzania, Zambia and Zimbabwe (2) (4).

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Physical Description

Morphology

The African hunting dogs' scientific name, Lycaon pictus, reflects the color of their pelage. Lycaon pictus literally means "painted or ornate wolf." The fur appears to be painted with brown, red, black, yellow and white areas. The pattern of colors is different on each animals coat, much like the stripes of zebras. The fur of L. pictus is short, with little or no underfur, and the blackish skin is sometimes visible where fur is sparse. Typically there is dark fur on the head and a white tip on the end of their bushy tail. They have large, rounded ears, a thin body, and long, muscular legs with four toes on each foot. The body length of Lycaon pictus is between 75 and 110 cm, the tail is between 30 and 40 cm long, and they range in weight from 18 to 36 kg. Males and females tend to be approximately the same size.

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Ecology

Habitat

Habitat and Ecology

African Wild Dogs are generalist predators, occupying a range of habitats including short-grass plains, semi-desert, bushy savannas and upland forest. While early studies in the Serengeti National Park, Tanzania, led to a belief that African Wild Dogs were primarily an open plains species, more recent data indicate that they reach their highest densities in thicker bush (e.g., Selous Game Reserve, Tanzania; Mana Pools National Park, Zimbabwe; and northern Botswana). Several relict populations occupy dense upland forest (e.g., Harenna Forest, Ethiopia; Malcolm and Sillero-Zubiri 2001). African Wild Dogs have been recorded in desert (Lhotse 1946) (although most desert populations are now extirpated), but not in lowland forest. It appears that their current distribution is limited primarily by human activities and the availability of prey, rather than by the loss of a specific habitat type.

African Wild Dogs mostly hunt medium-sized antelope. Whereas they weigh 2030 kg, their prey average around 50 kg, and may be as large as 200 kg. In most areas their principal prey are Impala (Aepyceros melampus), Greater Kudu (Tragelaphus strepsiceros), Thomson's Gazelle (Eudorcas thomsonii) and Common Wildebeest (Connochaetes taurinus). They will give chase of larger species, such as Common Eland (Tragelaphus oryx) and African Buffalo (Syncerus caffer), but rarely kill such prey. Small antelope, such as Dik-dik (Madoqua spp.), Steenbok (Raphicerus campestris) and Duiker (tribe Cephalophini) are important in some areas, and warthogs (Phacochoerus spp.) are also taken in some populations. African Wild Dogs also take very small prey such as hares, lizards and even eggs, but these make a very small contribution to their diet.

Generation lengthData on lifetime reproductive success of 19 alpha (breeding) females in western Zimbabwe indicate that 50% of reproductive output was achieved by age 5.5 years (SD 1.35, range 38; G.S.A. Rasmussen, unpubl. data). An alternative method, considers the average age of mothers of known litters, without the need for data on lifetime reproductive success. This method gives good agreement with the IUCN recommendations on calculating generation length, indicating a mean female breeding age of 5.7 years from the Zimbabwe dataset. Using this method, data from 18 litters born in Kenya to known-age mothers suggest a mean generation length of 5.0 years (R. Woodroffe, unpubl. data). Both studies suggest a minimum age at first breeding of approximately three years. Based on these data, for convenience we have estimated changes in African Wild Dog populations using a generation time of five years.

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African hunting dogs are found in grasslands, savannahs and open woodlands. They are widely distributed across the African plains and are not found in jungle areas. Their habitat also includes semi-desert to mountainous areas south of the Sahara Desert in Africa.

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The preferred habitats of the African Hunting Dog are savanna and open woodland. Once widespread, hunting dogs now survive in small fragmented populations in Namibia, Botswana, Mozambique,Zimbabwe, Swaziland, and the Transvaal.

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Trophic Strategy

African hunting dogs tend to prey on mammals that are about twice their weight. At times they will kill larger animals, and they will also take smaller prey individually. Some of the animals they prey on include small antelope such as impala (Aepyceros melampus) and bush duiker (Sylvicapra grimmia), and old, sick or injured larger animals such as wildebeest (genus Connochaetes) and zebra (genus Equus). On occasion some of the food they get from larger kills may be cached, though very often they never return to the cached food. For the most part Lycaon pictus does not eat plants or insects, except for small amounts of grass. Also African hunting dogs will never scavenge, no matter how fresh the kill is.

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Hunting dogs cooperatively hunt small antelopes such as impala and duikers, and old, sick or injured larger animals including wildebeest and zebra. They never scavenge, no matter how fresh the kill.

Highly social animals, hunting dogs form packs of up to 40 members, although average pack size is between 7 and 15. Pack members, led by a dominant female and male pair, cooperate in caring for and feeding the young and wounded or sick adults within their group.

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Lifespan, longevity, and ageing

Maximum longevity: 17 years (captivity) Observations: These animals have been estimated to live up to 17 years (Ronald Nowak 1999), though record longevity in captivity is only 15.1 years (Richard Weigl 2005). Further studies are necessary.

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Reproduction

Each African hunting dog pack has a dominant breeding pair. This pair can be identified by their increased tendency to urine mark. They are normally the only pair of pack members to mate and they tend to remain monogamous for life. Their life expectancy is approximately ten years. Generally the dominant pair prevents subordinates from breeding. Breeding suppression between females may often result in aggressive interactions. Occasionally a subordinate female is allowed to mate and rear young.

Mating System: monogamous ; cooperative breeder

Lycaon pictus reaches sexual maturity at approximately 12 to 18 months, though they usually do not mate until much later. The youngest recorded reproduction of a female was at 22 months old. Gestation is approximately ten weeks and pups are usually born between March and July. Litter sizes can vary considerably, from 2 to 20 pups. The smaller litter sizes have been recorded from animals in captivity. Breeding females gives birth to their litters in grass-lined burrows, usually an abandoned aardvark hole. The pups remain in the den with their mother for three to four weeks. Once the pups are brought out of the den they become the responsibility of the whole pack. Pups nurse from other females in the pack as well as from their mother. Weaning can occur as early as 5 weeks. The interval between litters is normally 12 to 14 months.

Breeding interval: Twelve to fourteen months

Breeding season: January to May

Range number of offspring: 2 to 20.

Average number of offspring: 8.

Range gestation period: 60 to 80 days.

Average gestation period: 72.4 days.

Range weaning age: 35 to 90 days.

Range age at sexual or reproductive maturity (female): 12 to 18 months.

African Wild Dogs have disappeared from much of their former range. Their population is currently estimated at approximately 6,600 adults in 39 subpopulations, of which only 1,400 are mature individuals. Population size is continuing to decline as a result of ongoing habitat fragmentation, conflict with human activities, and infectious disease. Given uncertainty surrounding population estimates, and the species tendency to population fluctuations, the largest subpopulations might well number <250 mature individuals, thereby warranting listing as Endangered under criterion C2a(i).

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Lycaon pictus is listed as endangered by the IUCN and the United States Endangered Species Act. Habitat loss and diseases that are spread by domestic animals jeopardize the remaining African hunting dog populations.

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Trends

Population

BackgroundAfrican Wild Dogs are rarely seen, even where they are relatively common, and it appears that populations have always existed at very low densities. Table 2 in the supplementary material (see below) presents estimates of the sizes of African Wild Dog subpopulations occupying all discrete areas of resident range identified by participants in the range-wide conservation planning process (IUCN SSC 2008, 2009, in prep.). These comprise a total of 39 distinct subpopulations estimated to range in size from two to 276 mature individuals.

Proportion of mature individualsEstimating the number of mature individuals is challenging, because African Wild Dogs are obligate cooperative breeders: within a pack, the alpha male and female are the parents of the majority of surviving pups (Girman et al. 1997). The IUCN Red List User Guidelines (IUCN 2010) define mature individuals as individuals known, estimated or inferred to be capable of reproduction, but do not specify the time period within which reproduction is considered possible. The User Guidelines go on to state in many taxa there is a pool of non-reproductive (e.g., suppressed) individuals that will quickly become reproductive if a mature individual dies. These individuals can be considered to be capable of reproduction.

In African Wild Dogs, a high proportion of individuals are indeed reproductively suppressed, but these animals do not always become reproductive quickly if an alpha individual dies. In a mature pack, most pack members are offspring of the alpha pair; for these animals, death of an alpha would usually not open up a breeding opportunity because no unrelated mates would be available within the pack. In our experience, death of an alpha often leads to disintegration of the pack, with no breeding until new packs are formed. Given these complexities, and in keeping with the spirit of capturing a snapshot of current conditions, we have chosen to define mature individuals as those considered capable of reproduction within the current breeding season. The number of mature individuals thus comprises the number of alpha males and females, and the number of sub-dominant (i.e. non-alpha) animals which breed successfully.

Table 1 in the supplementary material (see below) provides demographic data to allow the estimation of numbers of mature individuals (Nm) from the census population of adults and yearlings (Nc). Using these data, the number of alpha males (NaM) and females (NaF) would be estimated thus:

NaM = Nc x 0.55 x 0.176NaF = Nc x 0.45 x 0.215

Reassuringly, these equations return approximately equal estimates of the numbers of alpha males and alpha females.

No published estimate is available of the proportion of adults and yearlings which breed successfully as sub-dominants. However, Girman et al. (1997) report the proportions of surviving pups with sub-dominant mothers and/or fathers. Using a simple assumption that litter size was unrelated to social status, we therefore estimate the number of sub-dominant breeders (Nsub) as:

Nsub = (NaM x 0.10) + (NaF x 0.08)

The number of mature individuals is then calculated as

Nm = NaM + NaF + Nsub

We made exploratory calculations to investigate the effect of including the probability of currently-suppressed individuals surviving and breeding in subsequent years. These calculations became highly complex, and risked double-counting animals which might breed as sub-dominants in the current year and then attain alpha (dominant) status in later years. It is also important to note that, in the previous listing, the number of mature individuals was estimated simply as the number of alpha animals (i.e., NaM + NaF). These alternative ways of calculating numbers of mature individuals contribute to considerable uncertainty about African Wild Dog numbers.

Current population sizeTable 2 in the supplementary material (see below) presents estimates of the sizes of African Wild Dog subpopulations occupying all discrete areas of resident range identified by participants in the range-wide conservation planning process (IUCN SSC 2008, 2009, in prep.). These comprise a total of 39 distinct sub-populations estimated to range in size from two to 276 mature individuals. Few wild dog subpopulations have been systematically monitored, and these estimates are subject to considerable imprecision. This imprecision, combined with uncertainty about the calculation of mature individuals, and the species propensity for substantial population fluctuations, means that these population estimates should be viewed with great caution. Lack of precision has important consequences for Red List assessment; the two largest sub-populations (estimated at 268 and 276 mature individuals) are only slightly larger than one of the threshold sub-population sizes used by the Red List criteria (250 mature individuals: EN C2(i)). Had only alpha animals been considered mature individuals (as in the previous assessment), the sizes of these two largest sub-populations fall to 246 and 253 respectively. This uncertainty needs to be taken into account in comparing estimates of African Wild Dogs population sizes with IUCN Red List criteria.Change in population sizeData on African Wild Dogs past global population size are taken from Ginsberg and Woodroffe (1997). It is important to note, once again, that few of these population estimates are based on systematic monitoring, and all should be viewed with caution. Assessing changes in population size is complicated by the fact that a less complete dataset was available in 1997 than in 2012. As a result of these improved data, the global estimate of African Wild Dog population size is in fact higher for 2012 than for 1997. However, this difference reflects the greater area surveyed in 2012.

To try to overcome this problem, Table 3 in the supplementary material (see below) compares estimates of subpopulation size in 1997 (taken from Ginsberg and Woodroffe 1997) with estimates of African Wild Dog subpopulations from the same areas in 2012. Subpopulations known in 2012 but not known in 1997 are excluded. However, sub-populations known to be absent in 1997 (e.g. the managed metapopulation in South Africa, and the sub-population in Laikipia District, Kenya) are included in Table 3 . Hence, Table 3 represents our best assessment of a like-with-like comparison of African Wild Dog numbers in 1997 and 2012

Causes of declineThe causes of African Wild Dogs decline are reasonably well understood and include extreme sensitivity to habitat fragmentation as a consequence of wide-ranging behaviour, conflict with livestock and game farmers, accidental killing by people in snares and road accidents, and infectious disease. All of these causes are associated with human encroachment on African Wild Dog habitat and, as such, have not ceased and are unlikely to be reversible across the majority of the species historical range.Fluctuations in population sizePopulations of African Wild Dogs are prone to marked fluctuations at a variety of temporal and geographical scales which are likely to both increase extinction risks and undermine the precision of population estimates. At the local scale, a combination of high mortality, high fecundity, and dispersal by both sexes means that pack size fluctuates substantially over short periods (Figure 1 in the supplementary material), although fluctuation in numbers of mature individuals would be less dramatic. Because African Wild Dogs are seasonal breeders across most of their remaining geographic range, fluctuations may be synchronised across packs (see Figure 1 in the supplementary material).

The same demographic characteristics high mortality, high fecundity, and long-distance dispersal likewise lead to fluctuations at the population scale. This pattern is further exaggerated by the species susceptibility to infectious disease which can cause rapid die-offs. Local extinctions are not uncommon, and are often both rapid and unanticipated. Figure 2 in the supplementary material presents data from three relatively well-documented cases of local extinction involving small wild dog subpopulations affected by rabies.

Similar die-offs have been documented in larger African Wild Dog populations. For example, five of 12 study packs in Botswana (Alexander et al. 2010) and three of eight study packs in Kenya (Woodroffe 2011) have been reported as having died within short time periods during disease outbreaks. Although these relatively large study populations recovered, the majority of African Wild Dog sub-populations are estimated to comprise 20 mature individuals and could be severely compromised by outbreaks of this size.

For comparison, under good conditions African Wild Dog populations are also able to grow relatively quickly. African Wild dogs capacity for very long-range dispersal means that sub-populations sometimes reappear unexpectedly and grow rapidly; examples include natural recoveries in Samburu and Laikipia Districts, Kenya (Woodroffe 2011), Sav Valley Conservancy in Zimbabwe (Pole 2000), and the Serengeti ecosystem of Tanzania (Marsden et al. 2011). As an example of the speed of recovery, the subpopulation in Laikipia District, Kenya, grew from 0 in 1999 to 17 adults and yearlings in two packs by 2000, and by 2006 had increased 10-fold to 170 adults and yearlings (Woodroffe 2011).

On the basis of this evidence, we conclude that African Wild Dogs show substantial population fluctuations, but may not experience extreme fluctuations in sub-population size as outlined in the Red List guidelines. Nevertheless, the substantial fluctuations which do occur contribute to further uncertainty about sub-population sizes.

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African Hunting Dogs require large home ranges to support viable populations. In recent years, human encroachment has reduced and fragmented their habitat; domesticated dogs have introduced diseases such canine distemper and rabies with devastating consequences for hunting dog populations.

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Threats

The principal threat to African Wild Dogs is habitat fragmentation, which increases their contact with people and domestic animals, resulting in human-wildlife conflict and transmission of infectious disease. The important role played by human-induced mortality has two long-term implications. First, it makes it likely that, outside protected areas, African Wild Dogs may be unable to coexist with increasing human populations unless land use plans and other conservation actions are implemented. Second, African Wild Dog ranging behaviour leads to a very substantial "edge effect", even in large reserves. Simple geometry dictates that a reserve of 5,000 km contains no point more than 40 km from its borders a distance well within the range of distances travelled by a pack of African Wild Dogs in their usual ranging behaviour. Thus, from an African Wild Dog's perspective, a reserve of this size (fairly large by most standards) would be all edge. As human populations rise around reserve borders, the risks to African Wild Dogs venturing outside are also likely to increase. Under these conditions, only the very largest unfenced reserves will be able to provide any level of protection for African Wild Dogs. In South Africa, predator proof fencing around small reserves has proved reasonably effective at keeping dogs confined to the reserve, but such fencing is not 100% effective (Davies-Mostert et al. 2009) and is unlikely to be long-term beneficial for wildlife communities.

Even in large, well-protected reserves, or in stable populations remaining largely independent of protected areas (as in northern Botswana), African Wild Dogs live at low population densities. Predation by Lions, and perhaps competition with Spotted Hyaenas, contribute to keeping African Wild Dog numbers below the level that their prey base could support. Such low population density brings its own problems. The largest areas contain only relatively small wild dog populations; for example, the Selous Game Reserve, with an area of 43,000 km (about the size of Switzerland), is estimated to contain about 800 African Wild Dogs. Most reserves, and probably most African Wild Dog populations, are smaller. For example, the population in Niokolo-Koba National Park and buffer zones (about 25,000 km) is likely to be not more than 50100 dogs. Such small populations are vulnerable to extinction. "Catastrophic" events such as outbreaks of epidemic disease may drive them to extinction when larger populations have a greater probability of recovery such an event seems to have led to the local extinction of the small African Wild Dog population in the Serengeti ecosystem on the Kenya-Tanzania border. Problems of small population size will be exacerbated if, as seems likely, small populations occur in small reserves or habitat patches. As discussed above, animals inhabiting such areas suffer a strong "edge effect". Thus, small populations might be expected to suffer disproportionately high mortality as a result of their contact with humans and human activity.

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African wild dogs require large home ranges to support viable populations and recent habitat fragmentation has caused a population decline (4). Wild dogs traditionally have a reputation for attacking livestock, and despite this rarely occurring in practice, they are therefore often persecuted wherever they come into contact with humans (2) (4). In addition, road accidents and incidental snaring have recently become an important cause of mortality (2). Wild dogs are susceptible to disease; particularly those carried by domestic dogs such as canine distemper and rabies. A final threat that keeps wild dog populations low is competition and predation with the other large carnivores of the African savanna, such as lions and spotted hyaenas (6).

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Management

Conservation Actions

Conservation strategies have been developed for the species in all regions of AfricaSillero-Zubiriet al. 2004, Woodroffeet al. 1997, (IUCN SSC 2008, 2009, in prep.), and many range states have used these strategies as templates for their own national action plans (Department of Wildlife and National Parks 2008, Wildlife Service 2010). Although each regional strategy was developed independently through a separate participatory process, the three strategies have a similar structure, comprising objectives aimed at improving coexistence between people and African Wild Dogs, encouraging land use planning to maintain and expand wild dog populations, building capacity for wild dog conservation within range states, outreach to improve public perceptions of wild dogs at all levels of society, ensuring a policy framework compatible with wild dog conservation. These strategies are accessible at www.cheetahandwilddog.org.

Gaps in knowledgeSeveral pieces of information are needed to enable more effective conservation of African wild dogs. These include: 1) development of cost-effective methods for surveying wild dogs across large geographical scales; 2) surveys of wild dog distribution and status, particularly in Algeria, Angola, Central African Republic, Chad, Somalia, South Sudan, and Sudan; 3) development of locally-appropriate and effective means to reduce conflict between wild dogs and farmers; 4) establishing which techniques will be most effective and sustainable for protecting wild dogs from disease; and 5) determining the landscape features which facilitate (or prevent), wild dog movement over long distances and hence promote (or block) landscape connectivity.

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Conservation

The current population of African wild dogs is estimated to be less than 5,500 individuals (2). The majority of dogs are found in medium sized populations, which are extremely vulnerable to sudden environmental change. Due to their expansive home ranges, large areas of contiguous habitat are the key to the survival of this species and it is estimated that protected areas need to be greater than 10,000 square kilometres in order to prevent detrimental contact with local human settlements (2) (4). Consequently even most of the larger reserves cannot fully contain African wild dog packs, leading to clashes with humans (2). Preventing persecution through education is also a priority of the conservation action plan (2) (4), in an effort to preserve this most intriguing and unique canid.

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Wikipedia

African wild dog

The African wild dog (Lycaon pictus) is a canid native to Sub-Saharan Africa. It is the largest of its family in Africa,[2] and the only member of the genusLycaon, which is distinguished from Canis by its fewer toes and dentition, which is highly specialised for a hypercarnivorous diet.[3] It is classed as endangered by the IUCN, as it has disappeared from much of its original range. The current population has been estimated at roughly 39 subpopulations containing 6,600 adults, only 1,400 of which are fully grown. The decline of these populations is ongoing, due to habitat fragmentation, human persecution, and disease outbreaks.[1]

The African wild dog is a highly social animal, living in packs with separate dominance hierarchies for males and females.[4] Uniquely among social carnivores, it is the females rather than the males that disperse from the natal pack once sexually mature, and the young are allowed to feed first on carcasses. The species is a specialised diurnal hunter of antelopes, which it catches by chasing them to exhaustion.[5] Like other canids, it regurgitates food for its young, but this action is also extended to adults, to the point of being the bedrock of African wild dog social life.[6] It has few natural predators, though lions are a major source of mortality, and spotted hyenas are frequent kleptoparasites.[7]

The earliest possible written reference to the species comes Oppian, who wrote of the thoa, a hybrid between the wolf and panther which resembles the former in shape and the latter in colour. Solinus's Collectanea rerum memorabilium from the 3rd century AD describes a multicoloured wolf-like animal with a mane native to Ethiopia.[12]

The species was first described scientifically in 1820 by Coenraad Temminck, after having examined a specimen taken from the coast of Mozambique. He named the animal Hyaena picta, erroneously classifying it as a species of hyena. It was later recognised as a canid by Joshua Brookes in 1827, and renamed Lycaon tricolor. The root word of Lycaon is the Greek λυκαίος (lykaios), meaning 'wolf-like'. The specific epithet pictus (Latin for 'painted'), which derived from the original picta, was later returned to it, in conformity with the International Rules on Taxonomic Nomenclature.[13]

The English language has several names for Lycaon pictus, including painted lycaon,[12] African wild dog, Cape hunting dog,[8] and painted dog or painted wolf. The latter name is being promoted by some conservationists as a way of 're-branding' the species, as 'wild dog' has several negative connotations that could be detrimental to its image.[14] Nevertheless, the name 'African wild dog' is still widely used.[15]

The evolution of the African wild dog was once poorly understood, due to the scarcity of fossil finds. One proposed ancestral genus was Xenocyon, which lived throughout Eurasia, from Germany to Japan, as well as in Africa from the Early Pleistocene to the early Middle Pleistocene. The species X. falconeri shared the African wild dog's absent first metacarpal (dewclaw), though its dentition was still relatively unspecialised.[16] This connection was however rejected, as X. falconeri's missing metacarpal was a poor indication of phylogenetic closeness to the African wild dog, and the dentition was too different to imply ancestry. A more likely ancestral candidate is the Plio-PleistoceneL. sekowei of South Africa, on the basis of skull shape and tooth morphology, which shows the same adaptations to a hypercarnivorous diet as the modern species. L. sekowei had not yet lost the first metacarpal absent in L. pictus, and was more robust than the modern species, having 10% larger teeth.[3]

Fossil of Lycaon sekowei, a possible ancestor of the modern L. pictus.

Specimens inhabiting the Cape are characterised by the large amount of orange-yellow fur overlapping the black, the partially yellow backs of the ears, the mostly yellow underparts, and a number of whitish hairs on the throat mane. Those in Mozambique are distinguished by the almost equal development of yellow and black on both the upper and underparts of the body, as well as having less white fur than the Cape form.[20]

Similar to lupinus, but is smaller, has shorter and coarser fur, and has a weaker dentition. Its colour closely approaches that of the Cape form, with the yellow parts being buff rather than bright orange as is the case in lupinus.[20]

Nevertheless, although the species is genetically diverse, these subspecific designations are not universally accepted. It was once thought that East African and Southern African L. pictus populations were genetically distinct, based on a small number of samples. More recent studies with a larger number of samples showed that there has been extensive intermixing between East African and Southern African populations in the past. Some unique nuclear and mitochondrialalleles are found in Southern African and north-eastern African populations, with a transition zone encompassing Botswana, Zimbabwe and south-eastern Tanzania between the two. The West African L. pictus population may possess a unique haplotype, thus possibly constituting a truly distinct subspecies.[21]

The African wild dog is the bulkiest and most solidly built of African canids.[2] The species stands 60–75 cm (24–30 in) in shoulder height, and weighs 20–25 kg (44–55 lb) in East Africa and up to 30 kg (66 lb) in southern Africa.[5] Females are generally 3–7% smaller than males. Compared to members of the genus Canis, the African wild dog is comparatively lean and tall, with outsized ears and lacking dewclaws. The middle two toepads are usually fused. Its dentition also differs from that of Canis by the degeneration of the last lower molar, the narrowness of the canines, and proportionately large premolars, which are the largest relative to body size than any other carnivore other than hyenas.[4] The heel of the lower carnassial M1 is crested with a single blade-like cusp, which enhances the shearing capacity of the teeth and thus the speed at which prey can be consumed. This feature, termed "trenchant heel", is shared with two other canids: the Asian dhole and the South American bush dog.[8] The skull is relatively shorter and broader than that of other canids.[2]

The fur of the African wild dog differs significantly from that of other canids, consisting entirely of stiff bristle-hairs with no underfur.[2] It gradually loses its fur as it ages, with older specimens being almost naked. Colour variation is extreme, and may serve in visual identification, as African wild dogs can recognise each other at distances of 50–100 metres.[4] There is some geographic variation in coat colour, with north-east African specimens tending to be predominantly black with small white and yellow patches, while southern African ones are more brightly coloured, sporting a mix of brown, black and white coats.[8] Much of the species' coat patterning occurs on the trunk and legs. There is little variation in facial markings, with the muzzle being black, gradually shading into brown on the cheeks and forehead. A black line extends up the forehead, turning blackish-brown on the back of the ears. A few specimens sport a brown teardrop shaped mark below the eyes. The back of the head and neck are either brown or yellow. A white patch occasionally occurs behind the forelegs, with some specimens having completely white forelegs, chests and throats. The tail is usually white at the tip, black in the middle and brown at the base. Some specimens lack the white tip entirely, or may have black fur below the white tip. These coat patterns are asymmetrical, with the left side of the body often having different markings from that of the right.[4]

The African wild dog has very strong social bonds, stronger than those of sympatric lions and spotted hyenas, thus solitary living and hunting is extremely rare in the species.[22] It lives in permanent packs consisting of 2–27 adults and yearling pups. The average pack size in Kruger National Park and the Masai Mara is 4–5 adults, while packs in Moremi and Selous contain an average of 8–9. However, larger packs have been observed, and temporary aggregations of hundreds of individuals may have gathered in response to the seasonal migration of vast springbok herds in Southern Africa.[23] Males and females have separate dominance hierarchies, with the latter usually being led by the oldest female. Males may be led by the oldest male, but these can be supplanted by younger specimens, thus some packs may contain elderly former male pack leaders. The dominant pair typically monopolises breeding.[4] The species differs from most other social species by the fact that males remain in the natal pack, while females disperse (a pattern also found in primates like gorillas, chimpanzees and red colobuses). Furthermore, males in any given pack tend to outnumber females 3:1.[5] Dispersing females will join other packs and evict some of the resident females related to the other pack members, thus preventing inbreeding and allowing the evicted specimens to find new packs of their own and breed.[4] Males rarely disperse, and when they do, they are invariably rejected by other packs already containing males.[5] Although arguably the most social canid, the species lacks the elaborate facial expressions and body language found in the grey wolf, likely because of the African wild dog's less hierarchical social structure. Furthermore, while elaborate facial expressions are important for wolves in re-establishing bonds after long periods of separation from their family groups, they are not as necessary to African wild dogs, which remain together for much longer periods.[6]

African wild dog populations in East Africa appear to have no fixed breeding season, whereas those in Southern Africa usually breed during the April–July period.[22] During estrus, the female is closely accompanied by a single male, who keeps other members of the same sex at bay.[5] The copulatory tie characteristic of mating in most canids has been reported to be absent[24] or very brief (less than one minute)[25] in L. pictus, possibly an adaptation to the prevalence of larger predators in its environment.[26] The gestation period lasts 69–73 days, with the interval between each pregnancy being 12–14 months on average. The African wild dog produces more pups than any other canid, with litters containing around 6–16 pups, with an average of 10, thus indicating that a single female can produce enough young to form a new pack every year. Because the amount of food necessary to feed more than two litters would be impossible to acquire by the average pack, breeding is strictly limited to the dominant female, which may kill the pups of subordinates. After giving birth, the mother stays close to the pups in the den, while the rest of the pack hunts. She typically drives away pack members approaching the pups until the latter are old enough to eat solid food at 3–4 weeks of age. The pups leave the den at around the age of three weeks, and are suckled outside. The pups are weaned at the age of five weeks, at which point they are fed regurgitated meat by the other pack members. By seven weeks, the pups begin to take on an adult appearance, with noticeable lengthening in the legs, muzzle and ears. Once the pups reach the age of 8–10 weeks, the pack abandons the den, and the young follow the adults during hunts. The youngest pack members are permitted to eat first on kills, a privilege which ends once they become yearlings.[5]

The African wild dog is a specialised pack hunter of common medium-sized antelopes. Like the cheetah, it is the only primarily diurnal African large predator.[5]L. pictus hunts by approaching prey silently then chasing it in a pursuit clocking at 66 kmph for 10 to 60 minutes.[23] The average chase typically only goes as far as 2 km, during which time the prey animal, if large, is repeatedly bitten on the legs, belly and anus until it stops running, while smaller prey is simply pulled down and torn apart. L. pictus hunting strategies differ according to prey, with wildebeest being rushed at in order to panic the herd and isolate a vulnerable individual, whereas territorial antelope species, which defend themselves by running in wide circles, are captured by cutting off their escape routes. Medium-sized prey is often killed in 2–5 minutes, whereas larger prey like wildebeest may take half an hour to pull down. Male wild dogs usually perform the task of grabbing dangerous prey, such as warthogs, by the nose.[27] Small prey, like rodents, hares and birds are hunted singly, with dangerous prey like cane rats and porcupines being killed with a quick and well placed bite in order to avoid injury. Small prey is eaten entirely, while large animals are stripped of their meat and organs, with the skin, head, skeleton left intact.[22] The African wild dog is a fast eater, with a pack being able to consume a Thompson's gazelle in 15 minutes. In the wild, the species' consumption rate is of 1.2–5.9 kg per African wild dog a day, with one pack of 17–43 specimens in East Africa having been recorded to kill three animals per day on average.[15] Unlike most social predators, it will regurgitate food for adult, as well as young family members.[22] Pups old enough to eat solid food are given first priority at kills, eating even before the dominant pair; subordinate adult dogs help feed and protect the pups.[28] The African wild dogs is a highly successful hunter, with the majority of its chases ending in kills.[29]

The African wild dog is mostly found in savanna and arid zones, generally avoiding forested areas.[5] This preference is likely linked to the animal's hunting habits, which require open areas which do not obstruct vision or impede pursuit.[2] Nevertheless, it will travel through scrub, woodland and montane areas in pursuit of prey. However, forest-dwelling populations of African wild dogs have been identified, including one in the Harenna Forest, a wet montane forest up to 2400m in altitude in the Bale Mountains of Ethiopia.[30] There is at least one record of a pack being sighted on the summit of Mount Kilimanjaro.[5] In Zimbabwe, the species has been recorded at altitudes of 1,800 metres.[15]

Lions dominate African wild dogs, and are a major source of mortality for both adults and pups.[7] Population densities of African wild dogs are low in areas where lions are more abundant.[33] One pack reintroduced into Etosha National Park was destroyed by lions, and a population crash in lions in the Ngorongoro Crater during the 1960s resulted in an increase in African wild dog sightings, only for their numbers to decline once the lions recovered.[7] However, there are a few reported cases of old and wounded lions falling prey to African wild dogs.[34][35]

Spotted hyenas are important kleptoparasites,[7] and will follow packs of African wild dogs in order to appropriate their kills. They will typically inspect areas where African wild dogs have rested and eat any food remains they find. When approaching African wild dogs at a kill, solitary hyenas will approach cautiously and attempt to take off with a piece of meat unnoticed, though they may be mobbed in the attempt. When operating in groups, spotted hyenas are more successful in pirating African wild dog kills, though the latter's greater tendency to assist each other puts them at an advantage against spotted hyenas, who rarely work in unison. Cases of African wild dogs scavenging from spotted hyenas are rare. Although African wild dog packs can easily repel solitary hyenas, on the whole, the relationship between the two species is a one sided benefit for the hyenas,[36] with African wild dog densities being negatively correlated with high hyena populations.[37]

Although historically present, L. pictus is probably extinct, though may exist as a relict population in the south.[38]

As of 1997, the only recent reports come from the Teffedest Mountains. The species once occurred in the Mouydir Arah Mountains, but has disappeared, likely due to trapping and poisoning by Tuareg tribesmen. The last sighting in the Ahaggar National Park was in 1989.[38]

There was at least one unconfirmed sighting in 1992, and hunters living in the coastal areas of Western Sahara described a pack-hunting canid resembling L. pictus, though the identity of this animal is unconfirmed.[38]

The species is faring poorly in most of West Africa, with the only potentially viable population occurring in Senegal's Niokolo-Koba National Park. African wild dogs are occasionally sighted in other parts of Senegal, as well as in Guinea and Mali.[38]

The species may occur in Badiar National park, as the park is adjacent to Senegal's Niokolo-Koba National Park, where L. pictus does occur. The most recent reports of the species include a sighting in 1991 along the Sankarani River and the deaths of three cows in 1996 in the Ndama Fôret Clasée.[38]

The species is almost certainly extinct, having been the subject of an extermination campaign during the 1960s. Although legally protected, L. pictus specimens were still shot by game guards as recently as 1979. Even if still present, the species' chances of survival are still low, due to regular droughts and loss of natural prey.[38]

L. pictus may still be present in low numbers in Parc W, in the extreme north and the Sirba region.[38]

Although legally protected, there are no resident L. lycaon populations in Nigeria, though vagrants from neighbouring countries occasionally appear. Factors inhibiting the species' recovery include a lack of effective protection and the drastic reduction of its prey.[38]

L. pictus may still persist in low numbers in Gashaka Gumti National Park, which is fairly close to Cameroon's Faro National Park, where the species still occurs, though there were no sightings in 1982–1986. L. pictus is occasionally reported in Chingurmi-Duguma National Park, with the most recent sighting having occurred in 1995. It is likely extinct in Kainji National Park and Borgu Game Reserve, as poaching is intense and the species has not been sighted since the 1980s. It is also extinct in Yankari National Park, with the last sighting having taken place in 1978. One confirmed sighting of a lone individual occurred in 1991 in the Lame Burra Game Reserve.[38]

Although only partially protected, L. pictus has increased in number since the 1990s in and around Niokolo-Koba National Park, thus making Senegal the best hope for the species in West Africa.[38]

L. pictus is present in increasing numbers in and around Niokolo-Koba National Park. The population in the Park was estimated to number 50-100 specimens in 1997. This population is monitored and studied by the IUCN's Canid Specialist Group, in conjunction with Senegal's Licaone Fund. Elsewhere, L. pictus is rare or extinct.[38]

L. pictus may have once been present in the northern savannah-woodland areas, as natives there have names for the species, and some unconfirmed sightings were made in the 1980s. There may be a small population in Outamba-Kilimi National Park, though there has only ever been one unconfirmed sighting.[38]

The species is doing poorly in Central Africa, being extinct in Gabon, the Democratic Republic of Congo and the Republic of Congo. The only viable populations occur in the Central African Republic, Chad and especially in Cameroon.[38]

The status of L. pictus in Cameroon is uncertain, though three packs occur in the north of the country, thus making it the only possible refuge for the species in Central Africa, along with those present in CAR and southern Chad. Historically, most conservation efforts were directed to rainforest reserves, where L. pictus does not occur, though efforts in the 1990s sought to redress this. Nevertheless, attitudes towards the species remain negative, with 25 specimens having been killed by professional hunters in northern Cameroon in 1991–1992, with a government quota of 65 specimens during the December 1995 – May 1996 hunting season.[38]

The species is still regularly sighted in and around Faro National Park, where four packs were recorded in 1997. It is present in smaller numbers in Bénoué National Park, with several sightings having occurred in 1989 in the area between the two parks. L. pictus was sighted several times in and around Bouba Njida National Park in 1993.[38]

There are no recent reports of L. pictus in Chad, and their legal status is unknown. If the species does occur, then the southern part of the country may form an important link between Cameroonian and CAR L. pictus populations.[38]

L. pictus's range in East Africa is patchy, having been eradicated in Uganda and much of Kenya. A small population occupies an area encompassing southern Ethiopia, South Sudan, northern Kenya, and probably northern Uganda. The species may still occur in small numbers in southern Somalia, and is almost certainly extinct in Rwanda, Burundi, and Eritrea. Nevertheless, it remains somewhat numerous in southern Tanzania, particularly in the Selous Game Reserve and Mikumi National Park, both of which are occupied by what could be Africa's largest L. pictus population.[38]

L. pictus is rare in Ethiopia, despite total legal protection and the government's efforts at strengthening its network of protected areas. The species has been extirpated in three national parks, though it still occurs in the south of the country.[38]

The species was once occasionally recorded in and around Gambela National Park, though the last sighting occurred in 1987. It is frequently sighted in the Omo and Mago National Parks, with the most recent sighting in the former having occurred in 1995. Between 1992–1993, it was estimated that there were one or two packs in Omo and upt to five in Mago. It occasionally occurs in Bale Mountains National Park, though it is hampered by rabies and persecution by shepherds. Sporadic sightings have also occurred in the Awash and Nechisar National Parks. Three specimens were sighted in the Yabelo Wildlife Sanctuary in 1996. Outside protected areas, the species has been reported in Jijiga and Filtu.[38]

Although widespread, the species receives only partial legal protection, and primarily occurs in unprotected areas, with no high population densities. L. pictus numbers have declined, and it has become locally extinct in many areas, with only 15 packs occurring throughout the entire country as of 1997. Local attitudes towards it are poor, and it is frequently shot in livestock areas.[38]

Although legally protected, L. pictus is extinct in Rwanda, likely due to a disease outbreak. Modern Rwanda's overly high human population makes the country unsuitable for future L. pictus recolonisation, and a reintroduction project in 1989 was thwarted by the onset of the Rwandan Civil War.[38]

The species once occurred in high numbers in Akagera National Park, to the point of it being known as Le Parc aux Lycaons. A disease outbreak wiped out this population in 1983–1984.[38]

The ongoing Somali Civil War has made the outlook of L. pictus very poor in the country, with deforestation, poaching, drought, and over-grazing preventing the species from recovering, despite it being legally protected.[38]

The species may still be present in the north, though the last sighting occurred in 1982. It was once common in the Buloburde District before the late 1970s. A probably declining population may occur near the Jubba River. One pack was sighted in 1994 in Lag Badana National Park, which may be the best stronghold for the species in Somalia.[38]

As with all large carnivores, L. pictus populations fell drammatically during the Second Sudanese Civil War, though sightings have occurred in South Sudan.[38]

The species once occurred in the Sudd, though updates are lacking, and it is not afforded any legal protection in the area. It may be present in the Bangagai Game Reserve and Southern National Park. A pack was sighted in 1995 in Dinder National Park.[38]

Prospects in Tanzania are good for L. pictus, as the government imposed a moratorium on all hunting of the species, and it receives full legal protection. Although rare in the north, the south offers ideal habitat, as large tsetse fly populations prevent widespread human colonisation. The Selous Game Reserve and probably Ruaha National Park represent the best strongholds for the species in all of Africa.[38]

It is unlikely that Uganda has a resident L. pictus population, as the species was heavily persecuted after a 1955 directive to shoot it on sight. Vagrant specimens occasionally enter the country via Tanzania and South Sudan.[38]

A survey taken in 1982–1992 showed that the species was likely extirpated in Uganda, though sightings in some scattered areas may indicate that L. pictus is recolonising the country. Single individuals and small packs were sighted in Murchison Falls National Park, and were seen several times in the Northern Karamoja Controlled Hunting Area in 1994.[38]

Southern Africa contains numerous viable L. pictus populations, one of which encompasses northern Botswana, northeastern Namibia and western Zimbabwe. In South Africa, around 400 specimens occur in the country's Kruger National Park. Zambia holds two large populations, one in Kafue National Park, and another in the Luangwa Valley. However, the species is rare in Malawi, and probably extinct in Angola and Mozambique.[38]

Although L. pictus is legally protected, the Angolan Civil War prevented the collection of data, and there have been no reports of the species since 1990.[38]

The species was once found throughout Angola's protected areas, though it went into decline during the mid-1970s. It may still occur in the Cuando Cubango Province, where vagrants may arrive from Zambia and Namibia, though the population is probably unviable.[38]

The species' prospects in Botswana are hopeful, with the north of the country probably holding the largest L. pictus populations in all of Africa. Nevertheless, it receives only partial protection, and farmers are permitted to shoot it in defence of livestock.[38]

Although rare, L. pictus is legally protected, and may only be taken by government hunters and private citizens with ministerial permits. By the 1990s, it was regularly sighted in Kasungu National Park.[38]

The outlook of L. pictus in Mozambique is poor. The species underwent a rapid reduction in numbers after the Mozambican War of Independence in 1975, reaching the verge of extinction by 1986. Nevertheless, it regularly enters the country via Kruger National Park in neighbouring South Africa.[38]

L. pictus was once widely distributed in the remote and protected areas of the country, though it was declared extinct in western Manica, endangered in Tete and Zambezi, and extinct in Nampula. The species still occurred in the Rovuma and Lugenda River regions in 1986, and a pack with pups was sighted in Cahora Bassa in 1996.[38]

South Africa's L. pictus population is listed as 'specially protected' in the South African Red Data Book, and it has a stronghold in Kruger National Park, which held 350–400 specimens in the mid-1990s. There have been several attempts to reintroduce the species elsewhere, though only two of these attempts proved successful, and the resulting populations were not large enough to be viable.[38]

The species occurs in three regions: the Northern Cape, Kruger National Park, and KwaZulu-Natal. The Kruger population numbers at around 375–450 specimens, though they face pressure from lions and spotted hyenas, and are sometimes shot or snared outside Park boundaries. Six specimens were released into the Madikwe Game Reserve during the 1990s, though the reserve is too small to sustain a large population. In KwaZulu-Natal, the species is present in Hluhluwe–iMfolozi Park, where it was reintroduced during the early 1980s. This population has fluctuated since the reintroduction, and local attitudes towards it vary from hostile to favourable.[38]

Although once extensively persecuted, the species has total legal protection in Zambia, and can only be hunted after purchasing a costly licence from the Minister of Tourism. L. pictus remains widespread and occurs in most protected areas, which are large and hold suitable habitat and prey. Nevertheless, populations have declined since 1990.[38]

Zimbabwe holds viable L. pictus populations, which were estimated to consist of 310–430 individuals in 1985. The population increased during the 1990s, with a survey taken in 1990–1992 having estimated the population to be made up of 400–600 animals. The species is legally protected, and can only be hunted with a permit, which has only been given once between 1986–1992.[38]

Artistic depictions of African wild dogs are prominent on cosmetic palettes and other objects from Egypt's predynastic period, likely symbolising order over chaos, as well as the transition between the wild (represented by the golden jackal) and the domestic (represented by the dog). Predynastic hunters may have also identified with the African wild dog, as the Hunters Palette shows them wearing the animals' tails on their belts. By the dynastic period, African wild dog illustrations became much less represented, and the animal's symbolic role was largely taken over by the jackal.[9][10]

The African wild dog also plays a prominent role in the mythology of Southern Africa's San people. In one story, the animal is indirectly linked to the origin of death, as the hare is cursed by the moon to be forever hunted by African wild dogs after the former animal rebuffs the moon's promise to allow all living things to be reborn after death. Another story has the god Cagn taking revenge on the other gods by sending a group of men transformed into African wild dogs to attack them, though who won the battle is never revealed. The San of Botswana see the African wild dog as the ultimate hunter, and traditionally believe that shamans and medicine men can transform themselves into the animal. Some San hunters will smear African wild dog bodily fluids on their feet prior to a hunt, under the belief that doing so will gift them with the animal's boldness and agility. Nevertheless, the species does not figure prominently in San rock art, with the only notable example being a frieze in Mount Erongo showing a pack hunting two antelopes.[11]